Reuse & Permissions

It is not necessary to obtain permission to reuse this article or its components as it is available under the terms of the Creative Commons Attribution 4.0 International license. This license permits unrestricted use, distribution, and reproduction in any medium, provided attribution to the author(s) and the published article's title, journal citation, and DOI are maintained. Please note that some figures may have been included with permission from other third parties. It is your responsibility to obtain the proper permission from the rights holder directly for these figures.

Export citation

Export citation

Choose format for download:

Download Citation
  • Editors' Suggestion
  • Open Access

Phototactic Decision-Making by Microalgae

Shantanu Raikwar1, Adham Al-Kassem1, Nir S. Gov2,3, Adriana I. Pesci4, Raphaël Jeanneret1,*, and Raymond E. Goldstein4,†

  • *Contact author: raphael.jeanneret@phys.ens.fr
  • †Contact author: R.E.Goldstein@damtp.cam.ac.uk

Phys. Rev. Lett. 135, 228401 – Published 26 November, 2025

DOI: https://doi.org/10.1103/3fry-7tsw

Abstract

We study how simple eukaryotic organisms make decisions in response to competing stimuli in the context of phototaxis by the unicellular alga Chlamydomonas reinhardtii. While negatively phototactic cells swim directly away from a collimated light beam, when presented with two beams of adjustable intersection angle and intensities, we find that cells swim in a direction given by an intensity-weighted average of the two light propagation vectors. This geometrical law is a fixed point of an adaptive model of phototaxis and minimizes the average light intensity falling on the anterior pole of the cell. At large angular separations, subpopulations of cells swim away from one source or the other, or along the direction of the geometrical law, with some cells stochastically switching between the three directions. This behavior is shown to arise from a population-level distribution of photoreceptor locations that breaks front-back symmetry of photoreception.

View figure in article

Physics Subject Headings (PhySH)

Article Text

Supplemental Material

References (41)

  1. S. D. Fretwell and H. L. Lucas Jr., On territorial behavior and other factors influencing habitat distribution in birds, Acta Biotheoretica 19, 16 (1969).
  2. D. G. C. Harper, Competitive foraging in mallards: “Ideal free” ducks, Anim. Behav. 30, 575 (1982).
  3. J. Adler and W.-W. Tso, “Decision”-making in bacteria: Chemotactic response of Escherichia coli to conflicting stimuli, Science 184, 1292 (1974).
  4. L. R. Santos and A. G. Rosati, The evolutionary roots of human decision making, Annu. Rev. Psychol. 66, 321 (2015).
  5. C. R. Reid, S. Garnier, M. Beekman, and T. Latty, Information integration and multiattribute decision making in non-neuronal organisms, Anim. Behav. 100, 44 (2015).
  6. R. M. W. Chau, D. Bhaya, and K. C. Huang, Emergent phototactic responses of cyanobacteria under complex light regimes, mBio 8, e02330 (2017).
  7. M. Kim, Phototaxis of cyanobacteria under complex light environments, mBio 8, e00498-17 (2017).
  8. S. N. Menon, P. Varuni, and G. I. Menon, Information integration and collective motility in phototactic cyanobacteria, PLoS Comput. Biol. 16, e1007807 (2020).
  9. S. N. Menon, P. Varuni, F. Bunbury, D. Bhaya, and G. I. Menon, Phototaxis in Cyanobacteria: From mutants to models of collective behavior, mBio 12, e02398-21 (2021).
  10. M. Rivière and Y. Meroz, Plants sum and subtract stimuli over different timescales, Proc. Natl. Acad. Sci. U.S.A. 120, e2306655120 (2023).
  11. K. W. Foster and R. D. Smyth, Light antennas in phototactic algae, Microbiol. Rev. 44, 572 (1980).
  12. U. Rüffer and W. Nultsch, Flagellar photoresponses of Chlamydomonas cells held on micropipettes: I. Change in flagellar beat frequency, Cell Motil. Cytoskeleton 15, 162 (1990).
  13. U. Rüffer and W. Nultsch, Flagellar photoresponses of Chlamydomonas cells held on micropipettes: II. Change in flagellar beat pattern, Cell Motil. Cytoskeleton 18, 269 (1991).
  14. K. Schaller, R. David, and R. Uhl, How Chlamydomonas keeps track of the light once it has reached the right phototactic orientation, Biophys. J. 73, 1562 (1997).
  15. P. Hegemann, Vision in microalgae, Planta 203, 265 (1997).
  16. K. Josef, J. Saranak, and K. W. Foster, Ciliary behavior of a negatively phototactic Chlamydomonas reinhardtii, Cell Motil. Cytoskeleton 61, 97 (2005).
  17. K. Josef, J. Saranak, and K. W. Foster, Linear systems analysis of the ciliary steering behavior associated with negative-phototaxis in Chlamydomonas reinhardtii, Cell Motil. Cytoskeleton 63, 758 (2006).
  18. P. Hegemann, Algal sensory photoreceptors, Annu. Rev. Plant Biol. 59, 167 (2008).
  19. G. Jékely, Evolution of phototaxis, Phil. Trans. R. Soc. B 364, 2795 (2009).
  20. K. Drescher, R. E. Goldstein, and I. Tuval, Fidelity of adaptive phototaxis, Proc. Natl. Acad. Sci. U.S.A. 107, 11171 (2010).
  21. R. R. Bennett and R. Golestanian, A steering mechanism for phototaxis in Chlamydomonas, J. R. Soc. Interface 12, 20141164 (2015).
  22. H. de Maleprade, F. Moisy, T. Ishikawa, and R. E. Goldstein, Motility and phototaxis in Gonium, the simplest differentiated colonial alga, Phys. Rev. E 101, 022416 (2020).
  23. K. C. Leptos, M. Chioccioli, S. Furlan, A. I. Pesci, and R. E. Goldstein, Phototaxis of Chlamydomonas arises from a tuned adaptive photoresponse shared with multicellular Volvocine green algae, Phys. Rev. E 107, 014404 (2023).
  24. K. Yoshimura and R. Kamiya, The sensitivity of Chlamydomonas photoreceptor is optimized for the frequency of cell body rotation, Plant Cell Physiol. 42, 665 (2001).
  25. Z. Wang and A. C. H. Tsang, Intermediate light adaptation induces oscillatory phototaxis switching and pattern formation in Chlamydomonas, Proc. Natl. Acad. Sci. U.S.A. 122, e2425369122 (2025).
  26. R. L. Snyder and J. Dera, Wave-induced light-field fluctuations in the sea, J. Opt. Soc. Am. 60, 1072 (1970).
  27. J. Wei, M. R. Lewis, R. Van Dommelen, C. J. Zappa, and M. S. Twardowski, Wave-induced light field fluctuations in measured irradiance depth profiles: A wavelet analysis, J. Geophys. Res. Oceans 119, 1344 (2014).
  28. A. L’Homme, A. Lahlou, S. Bujaldon, T. Le Saux, B. Bailleul, N. Desprat, and R. Jeanneret, Light-induced phase separation with finite wavelength selection in photophobic microalgae, Phys. Rev. Lett. 135, 148401 (2025).
  29. A much earlier experimental study of the unicellular organism Euglena gracilis examined phototaxis in the presence of two lights, with adjustable intensities but a single angular separation, and found similar phenomenology to that reported here, described with an empirical fit to the data, D. P. Häder, M. Lebert, and M. R. Di Lena, New evidence for the mechanism of phototactic orientation of Euglena gracilis, Curr. Microbiol. 14, 157 (1986).
  30. V. H. Sridhar, L. Li, D. Gorbonos, M. Nagy, B. R. Schell, T. Sorochkin, N. S. Gov, and I. D. Couzin, The geometry of decision-making in individuals and collectives, Proc. Natl. Acad. Sci. U.S.A. 118, e2102157118 (2021).
  31. See Supplemental Material at http://link.aps.org/supplemental/10.1103/3fry-7tsw for further experimental results.
  32. J. C. Crocker and D. G. Grier, Methods of digital video microscopy for colloidal studies, J. Colloid Interface Sci. 179, 298 (1996).
  33. N. Isogai, R. Kamiya, and K. Yoshimura, Dominance between the two flagella during phototactic turning in Chlamydomonas, Zool. Sci. 17, 1261 (2000).
  34. C. R. Williams and M. A. Bees, Photo-gyrotactic bioconvection, J. Fluid Mech. 678, 41 (2011).
  35. L. de Andres-Bragado, C. Mazza, W. Senn, and S. G. Sprecher, Statistical modelling of navigational decisions based on intensity versus directionality in Drosophila larval phototaxis, Sci. Rep. 8, 11272 (2018).
  36. M. L. Zhu, K. J. Herrera, K. Vogt, and A. Bahl, Navigational strategies underlying phototaxis in Drosophila larvae, J. Exp. Biol. 224, jeb242428 (2021).
  37. N. R. Lebovitz and A. I. Pesci, Dynamic bifurcation in Hamiltonian systems with one degree of freedom, SIAM J. Appl. Math. 55, 1117 (1995).
  38. M. M. Moses, D. W. Morris, and W. Qin, Greener on the other side of the fence: Density-dependent habitat selection by a unicellular alga, Evol. Ecol. Res. 15, 1 (2013), https://www.scopus.com/inward/record.uri?eid=2-s2.0-84894144532&partnerID=40&md5=ee48fab54c523fc0cd948260afcad70c.
  39. J. Arrieta, A. Barreira, M. Chioccioli, M. Polin, and I. Tuval, Phototaxis beyond turning: Persistent accumulation and response acclimation of the microalga Chlamydomonas reinhardtii, Sci. Rep. 7, 3447 (2017).
  40. R. E. Goldstein, Green algae as model organisms for biological fluid dynamics, Annu. Rev. Fluid Mech. 47, 343 (2015).
  41. S. Raikwar, A. Al-Kassem, N. S. Gov, A. I. Pesci, R. Jeanneret, and R. E. Goldstein, Phototactic decision-making by microalgae, Zenodo, 2025, 10.5281/zenodo.15051634.

Outline

Information

Sign In to Your Journals Account

Filter

Filter

Article Lookup

Enter a citation